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P.S. 

Ro 

690 

B35 

1974 



Royal Ontario Museum 
Life Sciences 
Miscellaneous Publication 



Domestication of the Carp 
Cyprinus carpio L. 



Eugene K. Balon 







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Fig. 1 Coloration of Danube wild carp and reared offspring. Top: Danube wild carp, 
female. 49 cm standard length, from lesser Danube above Kolarovo. Slovakia. 
Centre: first generation of wild carp reared in ponds, cight-month-old fish, 15 
cm long. Bottom: seven-year-old, "dwarfed" Danube wild carp, 15 cm long 
(photo F. K. Balon). 



ROYAL ONTARIO MUSEUM 
LIFE SCIENCES 
MISCELLANEOUS PUBLICATION 



eugene k.balon Domestication of the Carp 
Cyprinus carpio L. 



ROM 



Presented to 



the Library 



of the 



Royal Ontario Museum 



by 



Dr. A.D. Tushingham 



Suggested citation: Roy.Ont.Mus. Life Sci. Misc. Pub. 
ISBN: 0-88854-147-3 
Publication date: 29 April 1974 



ROYAL ONTARIO MUSEUM 
PUBLICATIONS IN LIFE SCIENCES 

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life sciences contributions, a numbered series of original scientific publi- 
cations, including monographic works. 

LIFE sciences occasional pahers. a numbered series of original scientific 
publications, primarily short and usually of taxonomic 
significance. 

life sciences miscellaneous publications, an unnumbered series of 
publications of varied subject matter and format. 

All manuscripts considered for publication are subject to the scrutiny and 
editorial policies of the Life Sciences Editorial Board, and to review by persons 
outside the Museum staff who are authorities in the particular field involved. 

LIFE SCIENCES EDITORIAL BOARD 
Chairman: r. l. Peterson 
Editor: J. R. tamsitt 
Associate Editor: d. barr 
Associate Editor: e. j. crossman 



e. K. balon is Associate Professor. Department of Zoology, University of 
Guelph, and Research Associate. Department of Ichthyology and Herpetology, 
Royal Ontario Museum. 



price: $3.00 

©The Royal Ontario Museum, 1974 

100 Queen's Park. Toronto, Canada 

PRINTED AT THE BRYANT PRESS LIMITED 



Colin Bertram 
(Oryx, 1963. p.: 14) 



"Preservation by domestication in an alien en- 
viron/nent is far better than extinction in the 
natural habitat." 



I disagree 



In spite of commonly held views that domesti- 
cation preserves — it does not — it alters. 



H. Epstein 
(R. L. Smith's 

"'The Ecology of 
Man: . . .", 1972, p.91) 



"Domestication changed the life of the beast, the 
character of the animal, and its anatomy and 
physiology." 



The cover: Fishermen of the wild carp on the Danube River near Komarno one 
hundred years ago (reprinted from an etching in O. Herman, A Magyar halaszat 
konyve, Budapest, 1887). 



Contents 

Abstract, 1 

Resume. / 

Introduction^ 

Materials and Methods, 2 

Paleogeographical Distribution of Carp, 4 

Historical Evidence of Danubian Distribution, 5 

Taxonomy and Evolution, 7 

Dwarfed Wild Carp and their Viability, 8 

Probable Origin of Domestication, 16 

Roman Gourmets and First Carp in Captivity, 17 

Fasting Monks Rediscover the Carp, 1 9 

Rearing of Carp in Ponds and 

First Reproduction in Captivity, 20 

One More Origin of Domestication, 21 
Consequences of Domestication, 22 
Epilogue, 23 
Summary, 24 
Acknowledgments, 26 
Literature Cited, 27 
PE3IOME, 35 



'V\ 




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in 2012 with funding from 

Royal Ontario Museum 



http://archive.org/details/domesticationofcOObalo 



Domestication of the Carp 
Cyprinus carpio L. 



Abstract 

Historical, zoogeographical, morphological, and physiological 
information were utilized in explaining the origins and history 
of domestication of the carp. The wild carp ancestor apparently 
originated in central Asia and spread naturally east into China 
and west as far as the Danube River. Evidence suggests that 
the Romans first cultured carp collected from the Danube, and 
the tradition of the piscinae was continued in monasteries 
throughout the Middle Ages. A starvation experiment demon- 
strated the hardiness of Danube wild carp and confirmed its 
suitability for domestication. Domestication of the carp in 
China was first independent of similar activities in Europe. The 
strong evidence pointing to rheophilic Danube wild carp as 
ancestral to modern domestic varieties suggests the importance 
of preserving this natural population for further use. 



Resume 

Pour expliquer les origines et l'historique de la domestication 
de la carpe on a mis en oeuvre differentes sources de rensei- 
gnements; l'histoire. la zoogeographie, la morphologic et la 
physiologic. 

On trouve Tancetre de la carpe (a l'etat sauvage) apparem- 
ment d'abord en Asie centrale, puis graduellement vers Test 
jusqu'en Chine et vers l'ouest jusqu'au Danube. II y a tout 
lieu de croire que e'etaient les Romains qui firent les premiers 
l'elevagc de la carpe qu'ils prenaicnt du Danube, et on continua 
la tradition des piscinae dans les monasteres pendant le Moyen 
Age. On a tente de demontrer par des experiences avec la carpe 
sauvage du Danube, la resistance de celle-ci a un regime de 
jeune force, et ainsi on a meme prouve qu*on pourrait en faire 
parfaitement un elevage systematique. L'elevage de la carpe en 
Chine s'est developpe d*abord independemment de celui de 
l'Europc. Les recherches fakes nous ont fourni la quasi-certi- 
tude que la carpe sauvage reophile du Danube est bien 1'ancetre 
des differentes cspeccs moderaes d'elevage, et ainsi clles mct- 
tcnt meme au tout premier plan I'mterct a preserver cette popu- 
lation naturelle pour des usages ulterieurs. 



Introduction 

The wild carp (Cyprinn.s carpio carpio) is the predecessor of the domesti- 
cated or pond carp about whose vicissitudes we have less information than 
about the extinct auroch, the predecessor of cattle. An attempt is made, there- 
fore, to document its evolutionary and domestic history. 

During the past 400 years the domestic carp has been intensively cultured 
in Europe and introduced into many countries around the world. The origin, 
however, of this valuable commercial species has never been satisfactorily 
explained. The writing of this paper was stimulated by recent publications 
on the history of the carp in Europe (by Rudzihski, 1962, among others), 
Australia (Butcher, 1967), and North America (among others, McCrimmon, 
1968) that repeated old, unfounded ideas regarding the origin of this fish 
and also omitted details of its prehistory and early history (Vooren, 1972). 

Results of a number of studies (Balon and Misik, 1956; Misik, 1958; 
Misik and Tuca, 1965; Balon, 1957, 1958a, 1958b; Bastl, 1961 ; Rudzihski, 
1961 ) bearing on wild carp domestication are summarized here in an attempt 
to relate biological results to facts gleaned from zoogeography, history, and 
archaeology. As these data are meagre and their interpretation often ambigu- 
ous or subjective, the origin of domestic carp may not be solved definitively 
even in this essay. If my hypothesis, however, encourages more research and 
assists archaeology and social anthropology to interpret their newly discov- 
ered artifacts and data, the goal of this paper will have been accomplished. 

An earlier version of this study appeared in the mimeographed "Works of 
the Laboratory of Fishery Research in Bratislava*' (Balon, 1969). Presented 
here is a revised and amended version, with a new section on an experiment 
in starvation. The experiment was a result of an incidental activity, and 
though lacking details of a strictly scientific study, is worthy of record. 

Materials and Methods 

The first numerous wild carp in the piedmont zone of the Danube River were 
collected from a spawning school in May and June of 1955. Although I 
sampled individual specimens a year earlier (Balon and Misik, 1956), the 
intensified search resulted in finding the spawning school near the village of 
Kolarovo. These fish formed the main material for the age and growth study 
(Balon, 1957) and for the taxonomic revision (Misik, 1958). Although 
initially added to the collection of the Laboratory of Fishery Research (Slo- 
vak Agricultural Academy), most of them were later transferred to and made 
a part of the collection of the Slovak National Museum in Bratislava. Some 
specimens from successive spawning and developmental experiments (Balon. 
1958a), pond culture (Bastl, 1961) and from the experiment in starvation 
were also preserved and stored in both collections. These collections can now 
be considered major retainers of preserved rheophilic wild carp of the Danube 
River. 

The carp were kept in aquaria for an experiment in starvation (date of 
hatching — 31 May 1956). Individual specimens were successively selected 
for preservation or study of scales: first and second juveniles were preserved 



on 13 October 1956 (standard length [SL] 44 and 61 mm, weight [w] 2 and 
5 g respectively); third specimen, a mature female, was preserved 19 March 
1959 (SL 128 mm. w 56 g); from the fourth specimen scales were studied 
on 20 June 1958 (SL 129 mm, w 53 g) and the fish was preserved as a 
mature male on 19 March 1959 (SL 144 mm, w 67 g); from the fifth speci- 
men scales were studied on 20 June 1958 (when SL 120 mm, w 48 g) and 
the fish was preserved as a mature male on 9 December 1960 (when SL 134 
mm — the fish was unsuitable for most measurements because it jumped out 
of the aquarium and was found dehydrated); from the sixth and seventh 
specimen scales were studied on 13 October 1961 (SL 150 and 148 mm, 
w 69 and 56 g respectively) and on 20 June 1958 (when SL 138 and 146 
mm. w 63 and 80 g respectively), the sixth fish was preserved as a mature 
female on 26 April 1962 (SL 148 mm, w 47 g), the seventh fish as a mature 
female on 6 January 1963 (SL 148 mm, w 68 g). 

The relative growth indices used in Table 1 were explained in an earlier 
study (Balon, 1964, 1972). The identification of sex and usage of meristic 
and morphometric characters (Table 2) is defined in detail in Misik's (1958) 
study; the indices in Table 3 in Rudzinski's (1961) and Steffens' (1964) 
studies. 

Figures on the inside covers give some illustration of the extensive initial 
studies of the wild carp. They are reprinted here primarily for the benefit of 
the English language reader, and they explain the counts, measurements and 
usage of terms. 

INSIDE THE I RONT AND BACK COVERS 

Front cover: a-g some developmental stages of the wild carp (from Balon, 
1958a); a — embryo at time of hatching and its main respiratory organs — 
ducti Cuvieri in anterior part of the yolk sac; c — feeding larva 12 days old 
and 11 mm long with well developed (o) external lateral line sensors 
(cupulae); E — full grown larva 20 days old and 18 mm long; F — the appear- 
ance of first scales on a 15 mm long larva, and G — a 22 mm long juvenile. 
h-m some characters of an adult wild carp (from Misik, 1958) ; H — the type 
wild carp from the Danube River near Medvedovo (21.5.1954, Slovak 
National Museum #171 ), I — its first gill arch and J — pharyngeal bone; K — 
sketch indicating meristic characters; L, M — sketch indicating morphometric 
characters (numbers refer to characters listed in Table 2). 

Back cover: a — scaled morph, b — line or heavily scaled mirror morph, 
c — lesser scaled mirror morph. and d — scaleless or leather morph of the 
domestic carp (after Wunder, 1949); E — the type wild carp (from Balon, 
1967b); F — the rate of growth of wild carp (from Balon. 1957) ;g — the wild 
and H — the domestic carp (after Holcik and Hensel, 1972). 



Paleugeographical Distribution of Carp 

Studies proving the occurrence of the carp in Western Europe as early as the 
Tertiary are based primarily on scales found in preglacial freshwater strata 
in northern Germany (Nehring, 1883. after Zaunick. 1925) and at pile 
dwelling sites in Switzerland. Ruetimeyer's (1860) claim that he found 
scales of carp in remnants of pile dwellings was later contradicted by Forel 
(1904). Forel based his opinion on the information of Studer (in Zaunick. 
1925) who identified the scales as those of Abramis brama. Thus, not only 
is the identity of preglacial remnants doubtful, but their age is unconfirmed. 
There is no further true paleontological evidence. Consequently, the occur- 
rence of wild carp in preglacial Europe is conjectural. 

The most correct opinions seem to be those of authors (see in Misik, 
1958) who considered Asia Minor and the area of the Caspian Sea to be the 
origin of the wild carp. The primitive morph or subspecies of the wild carp, 
Cyprinus carpio anatolicus Hanko, 1924, with the greatest number of 
pharyngeal teeth, still occurs there. All morphs of the wild carp even today 
are highly adaptable and almost always react to changing environmental 
conditions by an abrupt modification of genotype (Dolzhenko, 1953; Bur- 
makin, 1956; and others). Therefore, all morphs of wild carp may be con- 
sidered to be evolutionarily young. That the primitive morph had started to 
occupy streams peripheral to western Asia where it originated as early as the 
late Pliocene is proved by remnants in "pontic" lacustrine strata (Borzenko, 
1926; Hanko, 1932; Banarescu, 1960). However, it probably did not survive 
the Pleistocene in that new area. Certain negative results support my hypothe- 
sis. Had the wild carp lived in the Danube refuge in the Pleistocene it would 
certainly have moved north to Scandinavia and the British Isles as conti- 
nental glaciers retreated before the end of the period of interfluvial connec- 
tions as did, for example, Abramis brama, Esox lucius, Rmilus rutilus, Scar- 
dinius erythrophthalmus, Tinea tinea, Blicca bjoerkna, and Cobitis taenia 
(Thiencmann, 1950). Carp also did not penetrate (e.g. via the Bering land 
bridge) to North America in the Pleistocene, at a time when this connection 
enabled Esox lucius to do so (Lindberg, 1962, Crossman and Harington, 
1970). Therefore the carp likely had not reached the Far East by the end of 
the Pliocene. Thus, the ancestors of recent carp probably evolved in the 
Caspian area and spread from there to western Europe and to China as late 
as the last postglacial period. 

Except for the Danube River, the natural occurrence of carp in waters of 
Europe as early as the beginning of the Christian era is not probable. Hence 
Ausonius (a.d. 310-393; 1933) did not mention carp in the fauna of the 
Rhine and Mosel rivers in the fourth century. Later records of sporadic 
occurrence of carp in rivers may be explained by individuals having escaped 
from ponds, though Dhigosz ( 1863-1887) did not mention the presence of 
carp in Polish waters even as late as the 1 5th century. 

The ancestor of contemporary wild carp evidently evolved in the area of 
the Caspian Sea at the end of the Pleistocene. Under conditions of the post- 
glacial thermal optimum some strains spread as far as the Black Sea area, the 
Aral system, into eastern Asia and appeared in the Danube River system 



about 8,000-10,000 years ago, that is, somewhat sooner than did the gold- 
fish. Carassius auratus gibelio, which today penetrates the Danube River 
(Hensel, 1971 ). The westward expansion of the range of that species was 
probably similar to that of the wild carp (Balon, 1962. 1963a). 

Jordan and Evermann (1896-1900, 1902) and Burns (1966) were thus 
mistaken to consider the carp to be an Asian species that was introduced in 
Europe before 1758 and was named after the Island of Cyprus, which they 
considered to be the centre of its distribution. Also in error was Butcher 
(1967) when stating. "'Whatever changes in the environment were brought 
about following the introduction of this fish into Europe have been lost 
apparently in antiquity." 

According to ancient stories and myths, the Great Schutt Island (in the 
upper part of the Middle Danube) was surrounded by a "great number of 
golden carp (Cyprinus auratus) that enabled even the poorest people to make 
a living; yes, there were times when the fishermen gave them away as gifts" 
(translated from Khin. 1930). Then why did Jordan (and Evermann, 1896- 
1902). Thienemann (1950). Maar (1960), Vooren (1972), and so many 
others believe that the carp reached Rome via Greece from China? Possibly 
even the carpio of Plinius (a.d. 24-79; 1635) was not an unknown seafish 
but the wild carp; then, of course, the specific name could be from the Greek 
root Karpos, meaning fruit. 

HISTORICAL EVIDENCE OF DANUBIAN DISTRIBUTION 

Natural occurrence of the wild carp in the Danube River is supported by the 
evidence of Lconhardt ( 1906), who however hypothesized that the wild carp 
arrived there from its northern distribution area before the formation of the 
continental pack-ice. Leonhardt (1906) assented that names used for the 
carp in local areas and the ultimate derivation of the scientific name can 
tell much about the historical distribution of that fish; he advanced a probable 
explanation of the generic and specific name of the carp. The generic name 
is from latinized Greek and the species name comes from the Celtic col- 
loquial name for the fish. "Kyprinos" or "Kyprianos" was the name given by 
Aristotle (384-322 B.C.; 1862) to this fish and was probably derived from 
"C\pris." a secondary name of the Goddess of Love, Aphrodite — perhaps 
because the high fertility of the carp was known even then. Later the name 
was latinized to "Cyprinus." probably by Pliny. Belon ( 1555) mentioned that 
he encountered the carp in the land of the Etolic Greeks, who called it 
""Kiprinos." 

The specific name, however, probably originated later and as "carpa" 
appeared for the first time in the works of Cassiodorus (a.d. 490-585; 1626) 
in the 6th century. The old Romans called it cyprinus, a name that spread 
with the fish throughout Europe. The lack of indigenous local names for this 
fish to the west and north of the Danube River may be considered to be 
further evidence that the name originated there with the appearance of the 
species. The Latin name of the fish is Celtic in origin and dates from the time 
when Celtic tribes inhabited the present eastern Austrian and Czechoslovak- 
Hungarian territory of the Danube River. Prom the ancient "charpho." 



"carfo" and "charofo" the name gradually changed to the present "carpo." 
Celts were certainly familiar with the carp at the time they settled in the 
Danube area, from where the name spread with them or with later introduc- 
tions of the fish. Everywhere the Celts lived, the root of the name "carp" oc- 
curs — "carpe" in France; "carpio" or "carpione" in Italy; "carp" in England; 
"carpe" in Spain; "Karpfen" in Germany; "karp" in Poland and Russia; and 
"kapr" and "Kapor" in Czechoslovakia. In the region of the Danube River, 
where it originated, however, this fish is called "ponty" (Hungarian), 
"sharan" (Serbian), "saran" (Bulgarian), "crap," "saran," "ciortan," 
"ciuciu" or "olocari" (Rumanian), "husgun" (Turkish), "korop," "sharan," 
"podrojek" (Ukrainian), "sazan" (on the Volga River), and "kalynshyr" 
(Kirghizia). 

Cassiodorus (a.d. 490-585; 1626) wrote about shipments of Danube 
carp to Ravenna in Italy for the table of King Theodorus, and his note may 
also serve as evidence of the Danubian origin of the western domestic carp. 
Consequently, there is no basis for later opinions which were copied without 
critical comment from one book to the next, that had the wild carp originate 
in China or eastern Asia and suggested that European pond-culture was, so to 
speak, the continuation of its earlier culture in China (Maar, 1960; Steffens, 
1967). These concepts were supported by some authors (Dubravius, 1547; 
von Hohberg, 1687) in the Middle Ages who wrote at a time when pond cul- 
ture was spreading but completely misplaced the origin of the fish. They 
found, correctly, that carp existed in European ponds before occurring in 
rivers but erroneously included the Danube among the rivers. According to 
von Hohberg ( 1687, p. 582) for example: 



Die Sce-Karpfen (odcr die in den 
Fliissen gcfangcn werden ) halt man 
auch fur besser (und wiewol sie ge- 
wohnlich in den Fliissen und Stro- 
men nich wohnen) geschiehct es 
doch vielmal (dass die abgebrochcn 
Teichc ihre Fische mit samt dem 
Wasser dahin uberlassen miissen) 
wie man an dem Teyalluss spuhren 
kan (dcr seine aus den Teichcn 
enpfangenc Karpffen der March) 
die March aber bei Toben der 
Donau mitt-heilet (daher auch 
zwischcn Prcssburg und Tbbcn vicl 
Karpffen in der Donau zu finden 
sind; sie gelangen zu grosscm Alter) 
wiewol sic in vicr und fiinf Jahren 
zu Speisc am dienlichsten. 



The lake carp (or that caught in 
rivers) is considered to be better, 
and although it ordinarily does not 
inhabit rivers and streams, it often 
finds its way into rivers with water 
from ponds with broken dams. 
Carp, released from ponds, are 
transferred first to the Dyje River 
and then to the Morava River, 
which joins the Danube at Dcvin. 
In the Danube River between Brat- 
islava and Devin occur many carp. 
Individuals reach a great age, al- 
though they are most edible when 
they are 4 to 5 years old. 



That individuals of the pond morph that escaped into rivers soon changed 
from a short, scaleless morph into an elongate, scaled morph similar to the 



wild carp supports von Hohberg's ( 1687) statement. A change in body shape 
is also known (Tchen. 1956) in varieties of the goldfish (Carassius auratus 
auratus). Although obser\ers in the Middle Ages considered the numerous 
carp at the mouth of the Morava River near Devin to have escaped from 
ponds, it was certainK the endemic, wild form of the carp that was abundant 
in that part of the river, which has a large flood zone needed for successful 
spawning of carp. 

Taxonomy and Evolution 

Recent native carp from the Danube River along the Czechoslovakia- 
Hungary border are an ancient morph and ecologically isolated from escaped 
or introduced pond carp ( Misik, 1 958 ) . But of all wild morphs of the carp — 
from the Danube River. Aral Sea. and central and eastern Asia — that in the 
Danube is nearest to the domestic carp (Balon, 1957, 1958a; Misik, 1958). 
In other words, we consider the wild carp of the Danube River to be not a 
form of pond carp which became feral after it was released into the river as is 
generally accepted, but as a true wild morph related to the domestic pond carp 
in a way similar to that of the ancestral auroch to cattle. In nature the wild 
carp does not hybridize with the domestic carp. Domestic carp are never 
found in schools of wild carp, and single, domestic carp caught within the 
spawning grounds of wild carp were never ready to spawn. Domestic carp 
which gradually change in rivers into a scaled, elongate morph similar to the 
wild carp, were normally easily recognized by a notch and hump posterior 
to the head; in native wild carp the transition between the top of the head 
and the back (nape) is slightly if at all marked. Numerous breeding experi- 
ments (Tuca. 1958; Bastl. 1961, 1962; Rudzinski, 1961, 1962; Misik and 
Tuca. 1965) proved the genetic distinctness of this morph. 

After a biometric study of wild Danube carp and after comparison with 
wild carp from other regions. Misik (1958) demonstrated that wild carp 
can be divided into three groups: 

1. European wild carp (Cyprinus carpio carpio) represented by the popula- 
tion of the Danube River system and designed as nominate subspecies; 

2. Eastern Asian wild carp (Cyprinus carpio haeinatopterus) from Siberia 
and China; 

3. Wild carp "from the Aral Sea and from other central Asian regions which 
in some ways are more closely related to the European ones, in others to east 
Asian carp, which at the same time are mutually substantially difTerent ,, 
(translated from Misik. 1958. p. 106). 

Concerning the third group Misik (loc. cit.) concluded (my translation) : 
"'Reasons for these convergent and divergent changes in morphological fea- 
tures of the central Asian carp — in my opinion — are to be sought in the geo- 
graphic conditions of the area, which may include specific peculiarities of the 
geography of both Europe and eastern Asia. It is particularly those specific 
characters that may show the orientation of the evolutionary divergence of 
central Asian carp. That is why they may be considered as intcr-digitating. 
allopatric populations, which — if they are geographically isolated from the 
subspecies C. c. haematopterus and C. c. carpio — can be considered as sepa- 



rate geographical morphs with numerous, ecological modifications." Misik's 
divisions, however, may be interpreted also as follows: 

The differentiation of the wild carp in the region of the Aral Sea and central 
Asia was influenced by an earlier geological age; if central Asia is considered 
to be the origin of the wild carp, descendants of that original strain had a 
longer time, the entire Pleistocene, to evolve in isolation as compared to 
carp that emigrated eastward and westward in the last postglacial period. 
Even though strains later integrated, more morphs may exist today in west- 
ern and central Asia than elsewhere. Regarding the low degree of differen- 
tiation, which nowhere reaches the species level, the wild carp is a geo- 
logically young species — dating at the most from the Pleistocene. From the 
western and central Asian area of first occurrence and origin of the wild 
carp, part of one strain later invaded the west, whereas part of another one 
has penetrated to the east. Consequently, Cyprinus carpio carpio and C. c. 
haematopterus exist today through their range as relatively poorly differen- 
tiated taxa. The degree of relationship between the diverse strains of central 
Asia and domestic stocks has yet to be established. 

Dwarfed Wild Carp and Their Viability 

Using wild adults selected on specially devised criteria (Misik and Tuca, 
1965), morpho-ecological characters were studied first (Balon, 1958a, 
1958b), and juveniles from the same spawn were used to evaluate the extent 
of changes following culture in southern (Bastl, 1961 ) and northern (Rud- 
zinski, 1961) European ponds (Slovakia-Poland). After developmental 
studies were terminated, some of these juveniles were maintained in aquaria 
in the laboratory and are discussed here. Though a more extensive study 
was originally planned, as a result of personal circumstances only a part of 
the data remained in my possession. In spite of this and the nine years 
elapsed since the termination of this experiment a record of the results may 
have some value. 

Seven fish, hatched from eggs collected after carp spawning in a pond on 
29 May 1956, were kept in small (30 1) aquaria. Each year, from 1956 
through 1963, some of the experimental fish were preserved. From the others, 
scales in the centre of body sides (key scales) were extracted, standard 
lengths and weights were recorded, and the fish were returned to the aquaria. 
The last fish was preserved after 7 years of continuous aquarium life. The 
average temperature was 21 C C (range 18-24°C), although in the winter of 
1958, as a result of a heating failure, the temperature was 10°C for several 
days. 

Throughout the experimental period the fish were fed mainly with Tubijex 
sp. at the rate of 1-2 g/week in the first 2 years and 3-4 g/week thereafter. 
Only on a few occasions was the Tubijex replaced by the same weight of 
scraps of beef or live Cladoccra. Although the diet was minimal to sustain 
life, in the first 2 years it was sufficient for some growth. The explanation 
could be that the ration provided, through that interval, some kind of nutrient 
more suitable for the developmental steps in question (Balon, 1971a) than 
for later steps, though even then the fish did not die as predicted by Vanccov 

8 



for such cases (Baton, 1960a ). During the 7 years of the experiment the fish 
behaved normally, and gonads developed each year in the last four years, 
although in the last five years the fish did not grow at all. These experiments 
demonstrated the ability of the wild carp to survive on small rations of sub- 
stitute food, the ability to survive starvation. (The term "starvation" is justi- 
fied by the minimal maintenance diet given and by the remarkable changes 
in body proportions of experimental fish.) As a consequence of starvation, 
however, the ossification of some skeletal elements of starved carp sets in 
earlier than in naturally developing carp (Balon, 1960b). The fish became 
asymmetrically "dwarfed" due to artificially induced achondroplasia-likc 
condition. 

Starvation affected growth of experimental fish, which never exceeded the 
average length and weight of the 1 -year-old fish from the natural habitat 
(Table 1 ). Nonetheless, both wild and experimental animals became sexually 
mature in the third growing season. 

Growth of experimental fish was more than six-fold less than that of wild 
fish in the Danube River. Furthermore, growth of experimental carp stopped 
in the fourth year. Differences in growth are reflected in relative indices 
(Table 1 ). The "index of average size" of the wild carp in the Danube was 
7 and only 2 in experimental carp; the "index of weight growth" among wild 
carp from a spawning school in the Danube River was 1052, whereas for 
first generation reared in ponds it was 962, and among experimental fish, 
196-1 

Sexually-mature gonads of experimental carp were first found in a male 
and female 34 months old preserved in March 1959. Ovaries of the female, 
which measured 128 mm (standard length), weighed 0.5 g (representing 
0.9% of the specimen's wet weight). The number of eggs was 3,950. 

Ovaries of a female that was preserved 3 years later (in April 1962; age 
5-f) weighed 2.8 g (left 1.56 g; right 1.16 g) and represented 5.8% of the 
wet fish weight. The number of eggs, estimated by the same method (gravi- 
metric), was 83,540. Ovaries of the last fish, a female that was 6-f years old 
and which was preserved in January 1963, were thin, compact strips adher- 
ing to the lateral and ventral sides of the intestine and contained large, single 
eggs that were irregularly distributed. New gonadal tissue had begun to de- 
velop around eggs from the previous year that had not yet been fully re- 
sorbed. a phenomenon that is not unexpected for a fish unable to spawn 
in an aquarium and forced to resorb the eggs for the last four years. Hence, 
in these experimental fish, age at first attainment of sexual maturity and num- 
ber of eggs produced are not related to size but to age. Did stress conditions 
of starvation produce changes leading to a new type of stock dynamic 
( Balon. 1 963b. p. 535 ) within the same generation? 

Starving did bring forth distinct changes in body proportions (Fig. 2) 
and in some meristic characters. Counts of fin rays of experimental fish are 
within the range of those of wild carp (Misik, 1958) inhabiting the Danube 
and those of the pond carp (Steffens, 1964). Scale counts and number of 
gillrakers are also within the range of normal variability. Pharyngeal tooth 
count and arrangement, however, are unusual (Table 2). Neither wild carp 



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formula 1.3-3.1. approx. x8. 



6. six-year-old female; tooth 



11 



TABLE 2. Morphometric and meristic characters of wild carp (Cyprinus 
carpio carpio) from the Danube River and of its "dwarfed" 
offspring from aquaria 



Meas- 






River 


Aquaria 


ure- 


Characters defined according to 


A 


' = 100 




N = 7 


ments* 


Misik, 1958 


X 


(range) 


X 


(range) 


(1-2) 


% OF STANDARD LENGTH: 










(1-3) 


Length of head 


24 


(21-27) 


30 


(29-33) 


(1-6) 


Length of snout 


9 


( 7-11) 


10 


( 9-10) 


(A-B) 


Length of barbel i 


1.7 


(0.3-3.1) 


4.2 


(3.3-5.2 


(C-D) 


Length of barbel n 


3.6 


(1.5-5.6) 


4.3 


(3.2-5.6) 


(6-11) 


Length of orbit 


3.3 


(1.9-4.7) 


7.4 


(6.7-9.0) 


(9-7) 


Interorbital width 


10 


( 8-12) 


10 


( 9-12) 


(11-3) 


Postorbital length of head 


12 


(10-14) 


14 


(14-15) 


(4-5) 


Depth of head 


20 


(17-23) 


22 


(19-25) 


(1-12) 


Predorsal length 


45 


(41-49) 


50 


(48-55) 


(1-13) 


Preventral length 


45 


(41-48) 


50 


(50-52) 


(1-14) 


Preanal length 


74 


(70-77) 


75 


(73-76) 


(12-15) 


Depth of body 


28 


(23-32) 


30 


(22-35) 


(12-26) 


Width of body 


17 


(11-23) 


15 


(13-18) 


(16-2) 


Length of caudal peduncle 


20 


(17-23) 


18 


(17-20) 


(18-19) 


Minimum depth of body 


12 


(11-14) 


12 


( 9-14) 


(20-13) 


Pectoral origin to pelvic base 


23 


(20-26) 


22 


(20-24) 


(13-14) 


Pelvic base to anal base 


30 


(26-33) 


26 


(26-28) 


(12-21) 


Length of base of dorsal fin 


40 


(36-45) 


36 


(34-38) 


(14-16) 


Length of base of anal fin 


9 


( 7-11) 


9 


( 7-10) 


(22-23) 


Length of upper lobe of caudal fin 


24 


(20-28) 


30 


(24-37) 


(24-25) 


Length of lower lobe of caudal fin 


24 


(20-28) 


30 


(25-38) 


(20-26) 


Length of longest pectoral ray 


19 


(15-22) 


21 


(19-25) 


(13-27) 


Length of longest pelvic ray 


17 


(14-20) 


20 


(18-22) 


(12-28) 


Length of longest dorsal fin ray 


17 


(13-20) 


20 


(18-22) 


(14-29) v 


Length of longest anal fin ray 


16 


(14-18) 


19 


(18-20) 


(1-3) 


% OF LENGTH OF HEAD: 










(1-6) 


Length of snout 


39 


(35-43) 


33 


(32-35) 


(A-B) 


Length of barbel i 


7 


( 3-12) 


14 


(11-18) 


(C-D) 


Length of barbel n 


15 


( 8-22) 


15 


(11-19) 


(10) 


Internasal width 


23 


(19-26) 


20 


(19-22) 


(6-11) 


Length of orbit 


14 


(12-17) 


24 


(22-28) 


(7-9) 


Interorbital width 


42 


(38-47) 


34 


(32-37) 


(11-3) 


Postorbital length of head 


51 


(47-54) 


46 


(45-49) 


(4-5) 


Depth of head 


84 


(74-95) 


73 


(66-79) 


(4) 


Width of head 


63 


(53-73) 


58 


(53-63) 


(2-16) 


% OF LENGTH OF CAUDAL 


PEDUNCLE: 






(16-17) 


Depth of caudal peduncle 


72 


(62-82) 


82 


(53-101) 


(17) 


Width of caudal peduncle 


41 


(30-52) 


33 


(24-46) 


(18-19) 


Minimum depth of body 


61 


(51-72) 


67 


(51-80) 


(13-20) 


",, OF PECTORAL ORIGIN TO 


PELVIC BASE: 






(20-26) 


Length of longest pectoral ray 


81 


(62-101) 


98 


(86-110) 


(13-14) 


% OF PELVIC BASE TO ANAL BASE: 






(13-27) 


Length of longest pelvic ray 


57 


(44-70) 


77 


(67-84) 



12 





Table 


2 (continued) 




Meas- 




River 


Aquaria 


ure- 


Characters defined according N = 100 


N = 7 


ments* 


to MiSik, 1958 


X (range) 


~x (range) 




FIN RAYS: 






(D) 


dorsal 


(ii) iii-iv 18-21 (22) 


in(18-19)20 


(A) 


anal 


(n) hi 4-5 


in (4) 5 


(C) 


caudal 


iv-viii 16-18 iv-viii 


vi-vii 17 (18) vi-viii 


(P) 


pectoral 


1(14) 15-18 (19) 


i(15) 16(17) 


(V) 


pelvic 


n 7-9 


I 8 (9) 


(K)lel't 


Total number of scales 


(34-36) 37-39 (40) 


(35) 36, 38 


right 


in lateral series 


(35-36) 37-39 (40) 


(35) 36 (37) 


above 


Number of scales in 


5-7 


5(6) 


lie low 


transverse series 


5-7 


5(6) 


(I) outsidi 


.' Number of gill-rakers 


(22) 23-27 (28) 


(22) 24, 26 (27) 


inside 


on the first gill arch 


(29) 30-34 (36) 


29(31) 


(J) 


Pharyngeal teeth 


1.1.3-3.1.1 


1.2.1-3.1 11.3-3.1 1 

1.2.1-1.2.1 



* See figures on inside front cover. 

nor domesticated pond carp had so many irregular teeth as experimental 
carp (Fig. 2), though domestic pond carp show similar irregularities in 15% 
more cases (Steffens, 1964) than wild carp. Dwarfed carp had considerably 
longer heads, a longer first pair of barbels, larger eyes, longer preventral 
distance, longer caudal fin lobes, and ventral fin rays (Table 2) than wild 
carp from natural habitats. Dwarf carp were also characterized by a slightly 
larger postorbital length of head, greater depth of head, greater predorsal 
length, greater depth of body, greater length of the longest pectoral fin, and 
longer ventral, dorsal, and anal fin rays in relation to the standard length. No 
characters are considerably smaller in the dwarf fish except interorbital width 
and postorbital length in relation to length of head. Length of snout, depth 
and width of head, and length of the dorsal fin base are somewhat smaller 
than those measurements in wild fish. 

Similar differences in body proportions between wild and domestic carp 
were found by Steffens ( 1964). Similar changes in morphometric characters 
were described by Wunder (1949) for starved carp and for starved Euro- 
pean bream (Abramis brama) by Liihmann and Mann (1957). In contrast, 
Rudzinski (1961 ) reported that wild Danube carp in ponds had a smaller 
head than a domestic carp of the type he studied. 

The size of the gape of the mouth, a character that Rudzinski ( 1961 ) and 
Steffens ( 1964) used so successfully to distinguish wild and domestic carps, 
was considerably smaller in the dwarfed fish than in the domestic or wild 
carp (Table 3). Rudzinski and Steffens considered the enlargement of the 
mouth in domesticated animals to be an adaptation to changes in feeding 
habits and also a probable result of artificial (man-induced) selectivity. 
Domestic carp selectively adapted to utilize supplementary food added to 
ponds grew better in ponds when artificial food was added. Is it possible 
that the small amount of food given in small particles to dwarf carp of my 



/.? 




Fig. 3 Scale of dwarfed wild carp no. 6 with 5 annuli; the second annulus (for 1958) 
is strongly intercepted, approx. xl6. 



experiment may have brought about the development of a small gape of 
mouth? The size of mouth would be an adaptive eharaeter in Kammerer's 
(1923) sense then, in spite of criticism of such concepts (Kocstler, 1971). 

The number of curves of the intestine in experimentally-dwarfed carp 
varied (4,7,4,4,5,6, respectively) as did other characters (Table 3). Adult 
domestic carp average six curves to the intestine (Klust, 1939). Hence not 
all dwarf fish developed fewer curves. Rudzinski ( 1961 ) found, and StefTens 
(1964) confirmed, that the intestine of wild carp was generally 15-25% 
shorter than that of domestic carp. Artificially "dwarfed" carp developed the 
shortest intestine which was less than 50% the length of that of the domestic 
carp and slightly more than 50% of that of the wild carp. Differences are 
even more emphasized if related to body weight: the ratio of gut length to 
body weight is 2.2 in domestic carp, 3.0 in wild carp, and 8.3 in dwarfed 
carp (Table 3). 

In spite of experimental conditions of minimal temperature fluctuation, 
the absence of seasonal changes, and constant feeding throughout the year. 



14 




Fig. 4 Scale of dwarfed carp no. 7, with absorbed dorsal and ventral edces, approx. 
x!8. 



scales of dwarfed carp developed regular annual marks (Fig. 3). The acci- 
dental drop in temperature in 1958 was indicated in the scales by a pro- 
nounced annulus. The shape of most key scales was regular but heavy 
absorption at the dorsal and ventral edges (Fig. 4) occurred in some col- 
lected toward the front or rear end of the body during the last two years. 
The first key scales were collected from the series above the lateral line, 
dorsal to the origin of the pelvic fin. True annuli were readily distinguished, 
but between them there were several confusing false annuli. I was, however, 
unable to relate scale growth to body growth, and values of body size back- 
calculated from scales for previous growth seasons do not make sense. 
Interception of annuli may be initiated by the intervals of life history and 
"the planetary yearly system" (Balon. 1971b. p. 99), but temperature or 
other environmental factors may strengthen the annulus interception (Seger- 
strale. 1932). 

The above data are recorded as a stimulus to interest, and in order to 
complete the description of the artificially dwarfed carp. The results of this 



15 



TABLE 3. Mouth-gape and intestine lengths indices for dwarfed, domestic 
and wild carp: o/l = 10 X mouth-gape (cm-')/standard length 
(cm), o/w = 10 X mouth-gape (cm-)/weight (g), o/lc = 10 
X mouth-gape (cm' J )/length of head (cm); gut/1 = length of 
intestine (tin) 1 cm of standard length, gut x w = length of 
intestine (cm)/ 10 g of weight. 



Index 




oil 


jw 


o/lc 


gut /I 


gut (w 




1 


0.10 


0.19 


0.32 


1.14 


21.22 




2 


0.38 


0.47 


1.17 


1.49 


18.09 


Dwarf wild carp 


3 


0.49 


0.11 


1.65 


0.67 


1.54 


(aquaria) 


4 


0.47 


0.10 


1.58 


0.84 


1.80 




6 


0.37 


0.12 


1.28 


1.10 


3.48 




7 


0.54 


0.12 


1.78 


1.77 


3.87 




X 


0.40 


0.18 


1.30 


1.17 


8.33 


Domestic carp 














(Rudzinski, 1961) 




1.91 










(Steffens, 1964) 














supplementary diet 




2.00 


0.08 


8.12 


2.64 


2.25 


natural diet 




2.30 


0.08 


8.12 






Wild carp 














(Rudzinski, 1961) 




1.14 










(Steffens, 1964) 














supplementary diet 




1.09 


0.10 


4.46 


2.11 


3.02 


natural diet 




1.24 


0.08 


5.27 







experiment extended conclusions of an earlier study (Balon, 1960b) beyond 
the early intervals of fish life. Karzinkin's (1935) experiments suggested 
that piscivorous fishes would be less capable of survival on substitute foods, 
although some species, at least, produce dwarfed morphs (Popova, 1967) 
if population survival is endangered. Some species that forage, however, 
may not only survive when starved but if later given an optimum diet over- 
take well-fed individuals (Krizenccky and Kfizenecka-Pulankova, 1953; 
Kuznccov, 1957). Great viability is a characteristic of wild carp, which toler- 
ate a wide range of environmental factors and arc consequently a unique 
telcost species — a most successful colonizer of the world. 

Probable Origin of Domestication 

The most western natural occurrence of rhcophilic wild carp in the Danube 
is at the mouth of the Morava (March) River. It was there, near Dcvin 
(Toben), that the well-known Amber Road crossed the Danube River. The 
Celtic settlements on the Danube River and the lower parts of its tributaries 
in southeastern Slovakia "existed until the middle of the first century a.d. and 
sometimes earlier (compare the settlement at Dcvin in the second half of 
the first century), for the Latin material culture often mixed with the Dacian" 
(translated from Pelikan, 1960). The carp was known to the Romans, who 
travelled along the Amber Road through territory where huge schools of 
wild carp probably spawned every year on Hooded meadows. Both facts indi- 

16 



cate that the Danube and Amber Road intersection is the site from which 
transfer of the carp to European waters outside the Danube area probably 
began. Other places are less probable, even though some were suggested. 
Movtchan ( 1966) even assumed that Romans transported carp to Italy from 
Fanagoria (the area of Kuban) and maintained them in ponds. Possibly, 
again, Dubravius (1547) was confused by the transfer of carp to brackish 
ponds and by its subsequent multiple spawning, which is, however, usual 
for carp in warm water (Busehkiel, 1932). Conseqently, Dubravius' inter- 
pretation of Pliny's (Plinius, a.d. 24-79; 1635) data concerning multiple 
spawning of carp in Italy was incorrect and distracted his attention from 
the true geographic origin of domestic carp. 

ROMAN GOURMETS AND FIRST CARP IN CAPTIVITY 

Although fish were captured in the Danube 10,000 years ago by the 
Cromagnons of Upper Paleolite (France), the first proof of fishing activity 
came from excavations of late Neolithic Stone Age (Clark, 1948). Childe 
(1929) assumed that ". . . important are the deer's horn harpoons from 
Vinca and Csoka (fig. 17), for they show how much the inhabitants of these 
sites appreciated the good fishing of the Danube." When the Romans arrived 
they encountered a well-developed fishing technique which was sufficiently 
documented in the first writings on fishing in the Danube River, scattered 
through the works of Strabo (63 B.C.; 1917-1932), Plinius (a.d. 24-79; 
1635). Aelianus (a.d. 170-235; 1858), and Ausonius (a.d. 310-393; 
1933). About a.d. 15 the Romans built a camp opposite the mouth of the 
Morava River on the site of a Celtic village that later became their biggest 
Pannonic town, Carnuntum, an important resting place on the Amber Road. 
Stone reliefs from that period frequently used the fish motif, which I had in 
two cases identified as a carp (e.g.. Fig. 5, Komarno City Museum). In the 
first century a.d. the XlVth and XVth Roman legions were stationed on the 
site of the present city of Bratislava, then called Peiso-Piso. Other garrisons 
and troops, living at sites along nearly the entire length of the Danube River, 
certainly fished for their livelihood as can be gathered from written remarks 
made by Roman authors spoken of earlier, as well as from works of Herod- 






Fig. 5 Stone relief with carp, partly spoiled by later sculptured head of a goat, from 
the Roman excavations at Komarom (Komarno City Museum). Photo E. K. 
Balon, 1966. 

17 



otus (485-425 B.C.; 1920-1924), Paterculus Velleius (19 b.c.-a.d. 31; 
1924), Tacitus (a.d. 55; 1836), and Cassiodorus (a.d. 490-585; 1626). 
The commanders of these garrisons were members of patrician families and 
as such were brought up on the Roman tradition of epicurianism. It must, 
therefore, have been a source of pride to bring back to Rome an as yet un- 
known delicacy. In Pliny's Historiae Naturalis, for example, are found such 
remarks as: 

Et in Danubio Mario extrahitur, And in the Danube River is fished 

porculo marino similimus . . . , hausen (Huso huso) resembling a 

sea dolphin . . . , 

and in Aelianus (loc. cit.) we find detailed descriptions of fishing through 
ice. As early as the first century B.C. Cicero's teacher of gastronomy, Sergius 
Orata, had devised special salt water reservoirs separated from the sea where 
he stored fish for the kitchen. These reservoirs ensured a permanent supply 
without regard to weather and success in fishing. According to Plinius (A.D. 
24-79; 1635) this method was adopted by Lucinius Muraena, who began 
storing freshwater fish. The patricians liked this manner of keeping their fish 
and competed in establishing such "piscinae," often spending enormous sums 
of money on them. Consul Lucullus (75 B.C.). whose reputation as a gourmet 
is well known, dug through a hill near Naples to bring water to his ponds, 
which were reputedly more costly than his villa. 

Although the original idea of the piscinae, as devised by Sergius Orata, 
was to store fish for gastronomic purposes, rearing fish later became a hobby. 
Another patrician, Hortensius, became as famous as Lucullus, but for the 
love of his ponds (Hortensius liked his eel so much that he wept when he 
found it dead . . .), and was even accused by Cicero of neglecting politics 
because of his fish. The Roman preferred sea fishes, as Varro (1 16-27 B.C.: 
1912) emphasized; freshwater ponds were apparently considered inferior 
and plebeian, but documented prejudice is at least proof of the existence of 
freshwater ponds. 

Now perhaps it can be understood why visiting senators, patricians, and 
plebeian soldiers not only tasted the fish during their temporary stay in the 
Danube region but tried to transfer them alive to piscinae at home, either as 
fish fanciers or as gourmet attractions for guests. Which of the largest, tastiest 
fishes of this region would have survived the rigours of primitive transport? 
Which could have lived in saline or brackish water after such transport? The 
wild carp is among the least sensitive of fish and can tolerate water with a 
low oxygen content so that it may be easily transported and afterwards be 
kept for weeks in small reservoirs (transportation in wet moss or other 
moisture-retaining materials is probable). Moreover, no special food is 
needed and it can endure long periods of starvation. Besides having palatable 
meat, carp has one very important ability — it lives in waters with salinities as 
great as 6,000 mg Cl/I (Nakamura, 1948; Johnson, 1954; Mark, 1966). 
Wild carp even live in the brackish waters of the delta of the Danube (Bana- 
rescu, 1964). 

The above, then, is how I envisage the Danube wild carp to have been 

IS 



transferred to Italy at the beginning of our era to begin life in piscinae. From 
there some escaped to local rivers and. alter the collapse of the Empire and 
the establishment of Christianity, were reared in monastery ponds. I do not 
exclude the possibility that wild carp endemic to the Danube River were later 
brought to other west European ponds. Apparently the messages of Roman 
and Middle Age scholars describing these early imports were passed on from 
generation to generation and probably accounted for the reestablishment of 
the carp-rearing tradition. In support of the above let me quote Cassiodorus 
(a.d. 490-585;"l622): 

. . . destinet carpam Danuvius: A ... from the Danube come carp 

Rheno veniat anchorago (...): and from the Rhine herring. To pro- 

sapori pisces de diversis finibus af- vide a variety of flavours, it is neces- 

ferantur. sic decet regem pascere, ut sary to have many fish from many 

a legatis gentium credatur paene countries. A king's reign should be 

omnia possidere. such as to indicate that he possesses 

everything. 

Apparently King Theodorus (a.d. 490-526) of Ravenna (Italy) ordered the 
transport of carp from the Danube to his country, thus imitating Roman 
tastes and continuing the still novel introductions. 

It was only 100 years ago that Dubisch brought Danube wild carp to the 
Upper Vistula River for hybridization with the local form of domestic carp 
(see Morcinek, 1909). In 1957 I repeated Dubisch's experiment by trans- 
ferring 1.000 yearling Danube wild carp to the ponds at Ochaby (experi- 
mental station of the Polish Academy of Sciences) (Rudzinsky, 1961 ). But 
as in experiments on the Danube, studies at the Ochaby pond proved (Lesz- 
czynska and Biniakowski. 1967) that wild carp are more suitable for stocking 
into natural waters than are domestic carp. 

FASTING MONKS REDISCOVER THE CARP 

Christianity introduced more than 100 fasting days a year, and the only meats 
that could be eaten during those days were crayfish, molluscs, and other cold- 
blooded animals such as fish (in some regions fowl and/or unborn rabbit 
embryos, laurices, were exceptions). As heavy punishment, which sometimes 
went as far as the death penalty, was meted for violations of proscriptions, a 
readily available supply of fish was important. Thus, fish became the only 
possible foods for monks during a substantial part of the year. Although fish 
were abundant in rivers and lakes, there were many days in the year when 
weather conditions, natural disasters, or wars made fishing impossible. 
Monks and missionaries, who had to follow fasting regulations, had difficulties 
finding fish (Leonhardt, 1906). Probably after monks settled and established 
a monastery, they thought about keeping a fish supply as they had in their 
southern homeland; they also remembered or read how suitable the carp was. 
The first monasteries were founded in the early 6th century (e.g., Monte 
Cassino Monastery was founded a.d. 529 ) . Later they gained land and farms, 
and conditions became favourable for the beginning of mass culture of fish 
that provided the monastery a ready supply of food for fast days. Monks, 
according to Leonhardt (1906). first reared local fishes: inland monasteries 

19 




Fig. 6 This illustration from the 18th century shows that monks, very much like the 
Romans, enjoyed fishing for carp in monastery ponds. (Courtesy of the 
Mansell Collection, London.) 

kept pike (Esox lucius), crucian carp (Carassius carassius) , bream (Abramis 
brama), and other species. But these species were difficult to keep in good 
condition in the primitive reservoirs. They knew that carp could be bred 
easily in these circumstances and introduced it. 

REARING OF CARP IN PONDS AND FIRST REPRODUCTION IN CAPTIVITY 

Carp then were first reared with other species of fish in a simple man-made 
rearing pond. Certainly, some unexpected spawning occurred already in 
Roman piscinae and also in monastery ponds; no organized reproduction, 
however, was recorded. Charlemagne (a.d. 768-814), the first Holy Roman 
Emperor, ordered his tenant farmers to maintain ponds and issued orders 
for their control. His orders, however, were concerned with protection against 
poaching, regulation of fishing and sale of fish, not with culture. Leonhardt 
(1906) claims that the lack of concern in carp reproduction was conditioned 
by an abundant fish fauna in local waters with which the landlords could 
regularly stock the ponds. In my opinion, Leonhardt (loc. cit.) incorrectly 
assumes the natural occurrence of the wild carp then in waters of southern 
and northern Europe. Moreover, archaeological findings of Slavonic settle- 
ments at the outset of the Polish Empire in the Ninth to Twelfth Centuries 
did not include carp, though they have produced remains of ide (Leuciscus 
idus), sturgeon (Acipenser sturio), chub (Leuciscus cephalus), tench (Tinea 
tinea), perch (Perca fluviatilis) , mud loach [Misgurnus fossilis), roach 
(Rutilus rutilus), beaked carp (Chondrostoma nasus). wels (Silurus glanis), 
pike (Esox lucius), eel (Ani>uilla anguilla), salmon (Sahno salar or Sabno 
trutta) and rapfen (Aspius aspius) (Perlbach, 1881 ). The first written evi- 
dence of carp in Polish territory is dated 1466 and originates from the area 



20 



of Kotomyje in the Black Sea Basin (Chmielewski, 1965). Later carp were 
kept in ponds as proved by the following words of M. Rej (a.d. 1505-1569) 
quoted by Gorzynski ( 1964) : 

... a wszystko to bardzo malem . . . and it is possible to obtain 

zachodem otrzymac mozna: bo sie- everything without trouble: release 

dem. ale dziewi^c karpi puscic, seven or nine carp, then the same 

takiez w druga (sadzawke) karas- number into another pond of cru- 

kow, ali ty i pieniazki i pozytek z cian carps, and you will benefit 

tego mice mozesz. from it as well as have money. 

But the pond culture of carp still cannot be connected with its pond repro- 
duction. 

Hildegarde ( 1089-1 170), mother superior of the Bavarian Convent of the 
Benedictine Order, mentioned in her recipes the preparation of carp (Koch, 
1925); Albertus Magnus (11937-1280; 1861) first wrote about breeding it 
in ponds. But generally not much information is available about the rearing 
of carp in the Middle Ages. It is as if the secret had been kept within the 
monastery walls. Gradually, however, complete articles, even simple studies 
about the rearing of the carp and its culture in ponds appeared (Dubravius, 
1547; Strumieriski, 1573; Strojnowski, 1609; von Hohberg, 1687), the rein- 
statement of rearing in the late Middle Ages is well known (Leonhardt, 1906; 
Steffens, 1958). Von Hohberg (1687) considered that the rearing methods 
of his time were superior to those used by the Romans and also that the 
Romans would surely have enjoyed the taste of the real contemporary carp. 
In the work of Dubravius (1547) numerous remarks about the beginning of 
Roman domestication of carp piece together the historical connections. 

ONE MORE ORIGIN OF DOMESTICATION 

Domestication of wild carp in China began independently, occurred five 
centuries earlier than in Europe (Leonhardt, 1906; Tamura, 1961; Hickling, 
1962), and probably involved the local subspecies Cyprinus carpio haema- 
topterus. The latest, anonymous Chinese study (the following quotations are 
from the Russian translation — Anonymous, 1961) on pond culture stated 
that "thanks to the creative efforts of the Chinese people for many genera- 
tions, breeding of the carp in this country has proceeded successfully for 
more than 2,000 .years. From China the breeding of this fish spread all over 
the world." Further, it was Fan Lio (Tao Tschshugun) who succeeded in 
spawning the carp and in growing the young to adulthood in a pond at the 
time of the "Spring and Autumn" and the "Fighting Dynasties" (in the 
eighth-third centuries B.C.). "From Asia the rearing of carp spread to 
Europe and later to America, Australia, and Africa. " There is no reason for 
the Chinese today to contradict an opinion that supports their national pride, 
particularly as most European authors supported the Chinese origin of the 
carp's domestication. The Chinese arc proud of their past isolation and inde- 
pendent cultural development. As China was virtually inaccessible until a 
century ago, would it not be improbable for the carp to have been brought 
from there in the first years of the Gregorian calendar or the beginning of the 
Middle Ages? An independent, second introduction of carp to Western 

21 



Europe probably occurred during the Middle Ages and is considered by some 
to have been from China (Rudzinski, 1962). Previously mentioned evidence 
favours the Danube as a source also of the later introductions. As a result of 
the introduction into Asia of European pond carp, however, the carp in ponds 
in some regions in eastern Asia may be mixed in origin (Buschkiel, 1933). 

Consequences of Domestication 

As early as the sixteenth century, races of carp were recognized, especially 
those morphs in which scales were few or absent. The domestic carp of 
western Europe was introduced into North America in 1831 for the first time, 
again in 1872, and frequently later (Hessel, 1881; Bartlett, 1901, 1905; 
Cole, 1905; Dymond, 1955; Atton, 1959; McCrimmon, 1968). The carp 
was introduced into Australia in 1860 (Butcher, 1962, 1967), and in 1896 
to the Cape of Africa (Jubb, 1967). According to my hypothesis all these 
carp originated from the rheophilic wild carp of the Danube River. While 
this hypothesis is geographically and historically attractive, in the absence of 
more evidence it remains still speculative. With this in mind, the gold carp 
(Cyprinus auratus) of the Schuykill and Massachusetts rivers (Forester, 
1850, quoted by McCrimmon, 1968) could be a scaled carp whose genotype 
was predominantly the wild morph and so selected a more lotic habitat than 
other carp in the shipment whose genes were primarily of more advanced 
domestic carp. (The most freshly caught Danubian wild carp that 1 saw were 
always a clear golden colour on the scaled region of their bodies.) 

In Europe the domestic carp became the most widely cultured and con- 
sumed fish, and pond culture gradually became one of the sophisticated 
branches of agriculture. The same species in North America, however, is 
considered to be undesirable, probably because of its unpalatable meat as 
compared to that of many local fishes and because of an adverse interaction 
with the aquatic habitat. Even in Australia "The European carp (Cyprinus 
carpio) has been proclaimed by legislation to be a 'noxious fish' and the 
keeping and/or release of this species is prohibited" (Butcher, 1967). 

Consequences of domestication of wild carp become apparent when the 
wild morph from the Danube is compared with any established domestic 
morph. Subsequent to rediscovery of rheophilic wild carp in the Czecho- 
slovak-Hungarian section of the Danube (Balon and Misik, 1956) and 
initial studies on it (Balon, 1957, 1958a; Misik, 1958), morphomctric and 
growth indices were compared with those of the domestic populations (Tuca, 
1958; Steffens, 1959; Bastl, 1961, 1962; Rudzinski, 1961; Chytra ct al., 
1961 ; and Misik and Tuca, 1965). Some morphological differences between 
wild and domestic carp were stressed by Rudzinski (1961) and Steffens 
( 1 964 ) . It will be sufficient to mention some of the results of the latter study : 

The body of the domestic carp, which is nearly always much deeper than 
that of the oblong, cylindrical body of the wild carp, appears to have more 
flesh, but the calculated ratio of muscle in both morphs is the same. Without 
regarding changes "in proportions, dressed weight of individual domesticated 
carp did not increase even though its greater growth rate produces more 
absolute meat in a given period of time. Probably the faster growth of domcs- 

22 



tic carp can be correlated with the larger mouth and longer intestines acquired 
by the pond form following its adaptation to complementary nutrition and 
perhaps also by man's selection. 

The greater strength, mobility, and viability of wild carp are emphasized 
by physiological characteristics. The wild carp has 189c to 19% more eryth- 
rocytes and haemoglobin than docs the domestic carp. Blood sugar level is 
16%— 26% higher. The wild carp has a much lower water content in muscles 
and liver than does the domestic carp. Furthermore the wild carp has a 
greater content of fat in individual organs, of glycogen in the liver, and of 
vitamin A in the intestines, eyes, and liver. Consequently the taste of the wild 
carp is better because the flesh is juicier. The wild carp is more mobile, 
stronger, and nimbler because its muscles are better supplied with blood 
ensuring a better supply of nutrients and elimination of waste; muscles do 
not fatigue as quickly as do those of the domestic carp, enabling wild carp to 
overcome the river current. The same improves the taste of the flesh of the 
wild hare and the wild rabbit, said to be more juicy than the domestic rabbit 
(Volf, 1965). 

Epilogue The impressive number of these 

that are 'not known in the wild state' 
emphasizes the drastic nature of the 
morphological changes that domes- 
tication has so often brought about 
in the transition from wild ancestor 
to cultivate crop. 

David R. Harris (1967, p. 91) 

Perhaps the unpalatable taste of carp meat in North America is a result of 
the carp's origin. Nearly all carp that live in natural waters here are descend- 
ants of domestic carp which escaped into natural habitats. Limited ability to 
utilize natural food and poor growth may have resulted in unpalatable flesh 
in feral domestic carp. European domestic cattle, when left to become wild, 
are known to remain in poor condition and never revert to a condition 
similar to that of the strong, wild, ancestral auroch (Bos primigenius) (see 
Talbot et al., 1965; Taylor. 1970. 1972). Perhaps the carp fall to feral con- 
dition parallelled that of the cattle. If this is the case it is not surprising that 
in North America carp "have never been wholeheartedly accepted by the 
general public as an item of food" (Crossman, 1 969. p. 1 45 ) . 

The need for fasting food may have been the main reason for rediscovery 
of the carp in the Middle Ages, but not everywhere has the hedonistic 
approach of the Romans to rearing of the carp given way to pure necessity; 
there is some evidence (Fig. 6) of monks enjoying fishing for carp (Burton 
and Burton. 1 968 ) . Nevertheless, the need for a larger variety of foods during 
fasts undoubtedly played an important role in bringing about the tradition of 
carp eating among some European Christians. Though related to fasting, this 
tradition was eventually also embraced by some European Jews who made 
the carp their Sabbath meal. 

23 



That the history of the domestication of the carp is generally similar to 
that of the European rabbit (Oryctolagus cuniculus) may be of some interest 
as peripheral support of my hypothesis on carp origin. Varro (1 16-27 B.C.; 
1912) wrote that Romans brought rabbits from Spain and bred them in 
special enclosures called leporaria to ensure a fresh supply of meat at all 
times. Unborn rabbits collected from pregnant females were prepared as a 
special dish called laurices. At the beginning of the Middle Ages rabbits were 
a popular fare in western European monasteries, and laurices were eaten 
even during fasting periods. (It was in the corridors and paved courts of the 
monasteries that the rabbits began giving birth to their young above ground 
instead of in burrows. The young gradually became domesticated and accus- 
tomed to people.) In the middle of the sixteenth century several colour 
morphs of rabbits were known. An albino rabbit is shown in Titian's 
"Madonna" (1530) at the Louvre (Valcanover, 1960). Around 550 B.C. 
the Chinese philosopher Confucius (1898) suggested to poor farmers that 
they keep some rodents similar to rabbits (Volf, 1965). Unlike speculation 
on the origin of the carp, the origin of the domestic rabbit was not assumed 
to be in China. Otherwise the similarity of the domestic history of both 
animals is striking. 

Wild rabbits are overabundant in many countries of the world and conse- 
quently special measures are taken to control them. Conversely, rheophilic 
wild carp in Europe today is in danger of extinction. Regulation of the flow 
of rivers and construction of waterworks arc destroying remaining spawning 
areas. In the Danube River, between the mouths of the Morava and Hron 
Rivers, spawning schools that were plentiful as recently as 15 years ago have 
greatly decreased. At a decreased population density the wild carp may lose 
its habitat to the domestic carp, which is repeatedly stocked, purposefully 
and accidentally, into rivers. Competitive replacement may be accelerated if 
planned hydroelectric plants and waterways are introduced (Balon, 1967a, 
1967b). 

Scientifically directed selection would have a better chance of improving 
cattle production and quality if genes of the ancestral auroch were still 
available. The wild carp may be compared to the extinct auroch when 
consequences of its possible extinction are contemplated. In 1965 Slovakia 
included spawning schools of wild carp in the official list of animals protected 
by law (Randik, 1967). Even so, because of the rapid deterioration of the 
environment, extinction is the probable fate of this species. Protection of the 
original stock of wild carp in the foothills of the Danube River is therefore 
an important task for mankind. It was probably here that the most useful 
domestic fish of the world originated. 

Summary 

If preglacial remnants of the carp from pile dwellings are rejected as incor- 
rectly dated, it is postulated that the wild carp originated in central Asia and 
spread naturally east into China and adjacent regions and west as far as the 
Danube River in the last postglacial period. That this did not happen earlier 

24 



is suggested by the fact that in the Pleistocene, after the retreat of the conti- 
nental glaciers, C. carpio was not among the species which penetrated either 
Scandinavia, the British Isles, or North America. Had it occurred at that time 
in eastern Asia it probably would have invaded North America together with 
the pike and other freshwater fishes via the Bering land bridge. 

The natural occurrence of the wild carp in the Danube River is inferred 
from the different names given to the carp by the local inhabitants and from 
the use of the same Celtic name outside the western limit of its natural range; 
further evidence comes from medieval chronicles concerning the occurrence 
of this fish, and from contemporary studies of the rheophilic wild form of the 
carp in the Danube River. The ancient differentiation of wild carp in central 
Asia and presence of undifferentiated isolated single morphs in the eastern 
and western regions where it occurs suggest again that its place of origin must 
be sought in central Asia. 

A separate section is devoted to an experiment testing the ability of Danube 
River wild carp to survive in starvation condition; this demonstrates the 
hardiness of the animal and explains why it was domesticated for human 
consumption. 

Finally, from written notes of the Romans and excavated art objects de- 
picting the carp, the theory is presented that the first carp brought to Roman 
piscinae came from places where the Amber Road — the most travelled thor- 
oughfare of those times — crossed the Danube River. These localities, regu- 
larly inundated meadows where schools of carp spawned, occurred at the 
mouth of the Morava River, near ancient Carnuntum and the later Devin 
Castle. Patrician commanders of the Roman legions stationed there brought 
the carp to Rome as an epicurian delight, for it was one of the most hardy 
species of fish and survived primitive techniques of transport and rearing. 
After the decline and fall of the Roman Empire, the rearing techniques of 
carp in Roman piscinae were not forgotten. Christian missionaries and monks 
needed ready reserves of fish for their fasting days. From Latin writings and 
from traditions of their southern homelands they knew that the carp was 
appropriate for such purposes and looked after their supply as soon as they 
settled somewhere. After the Middle Ages breeding had become routine, not 
only in monasteries but also on private estates. 

The history of the domestication of the carp in Europe that began with 
wild carp from the Danube River (Cyprinus carpio carpio) is divided into 
five periods: 1. First introduction by the Romans west of the Danube River 
in the first to fourth century a.d.; 2. sporadic western introduction and rear- 
ing in the fifth and sixth centuries; 3. beginning of rearing on a mass scale, 
with some renewed introductions, and initial attempts to breed it in greater 
quantities in specially-built ponds in the seventh to thirteenth centuries; 
4. beginning of breeding, mass culture, and fortuitous selection in the four- 
teenth to sixteenth centuries; 5. intensification of breeding, purposeful selec- 
tion, and introductions into North America. Australia, the Far East, and 
Africa from the seventeenth century. In China domestication of the carp, 
probably from local subspecies C.c. haematopterus, was begun five cen- 
turies earlier but independently of similar efforts in Europe. 

25 



The most important features by which the wild carp from the Danube 
River differ from the present domesticated forms are stressed and reference 
is made to a striking similarity with the domestication of the rabbit. Attention 
is drawn to the necessity of preserving the last wild schools of this fish in the 
foothill-zone of the Danube River and of protecting the wild predecessor of 
the domestic carp and its habitat. Most important, a repetition of the fate of 
the extinct auroch, the ancestor of cattle, should not be permitted. As with 
cattle, domestic carp in wild conditions becomes an animal of poor food 
quality, which is perhaps the reason why the carp deteriorated so after its 
transatlantic introductions. 

Acknowledgments 

Above all, I owe an outstanding debt of gratitude to V. Misik, V. Tuca, I. 
Bastl (Slovak Agricultural Academy, Bratislava) and E. Rudziriski (Polish 
Academy of Sciences, Cracow) for their original studies of the Danube wild 
carp that were consistent with my ideas and which led to the formulation of 
the theorem here presented. I thank J. Holcik (Slovak National Museum, 
Bratislava), O. Oliva (Charles University. Prague) and A. G. Coche 
(UNDP/FAO, Chilanga), who pointed out errors or inadequacies in the 
first draft, and E. J. Crossman (Royal Ontario Museum, Toronto) for read- 
ing, correcting and criticizing the final draft. 



26 



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1953 K biologii akklimatizirovannogo sazana v ozerakh zapadnoi Sibiri [On 
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29 



GORZYNSKI, S. 

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1932 Ursprung und Verbreitung der Fischfauna Ungarns. Arch. Hydrobiol., 
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HENSEL, K. 

1971 Some notes on the systematic status of Carassius auratus gibelio (Bloch. 
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1921- Herodotus, with an English translation by A. D. Godley. 4 vols. London. 
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1881 The carp and its culture in rivers and lakes; and its introduction into 
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1962 Fish culture. London, Faber and Faber. 295 pp. 

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1687 Georgica Curiosa oder Adeliches Land und Feld Leben. it Theil. Niirnberg. 
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1972 Ichtyologicka prirucka [Ichthyological Handbook]. Bratislava. Obzor. 
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1954 Preliminary experiments on fish culture in brackish-water ponds. Progve. 
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1967 Freshwater fishes of Southern Africa. Capetown, A. A. Balkema. 248 pp. 

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1935 K poznaniyu rybnoy produktivnosti vodojemov. Soobshtclienye IV. Usvoenie 
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30 



KIIIN, A. 

1930 Vyzy no Vel'kom Zitnom ostrove a ich lovenie [Hausen on Great Shutt 
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1939 Uber Entwicklung, Ban und Funktion des Darmes beim karpfen (Cyprinus 
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1925 Die Geschichte der Binnenfischcrci von Mitteleuropa. In Handbuch der 
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1971 The Case of the Midwife Toad. New York. Random House. 187 pp. 

KRIZENECKY. J. AND A. KR1ZENECKA-PI I \NKOV\ 

1953 K otazce rustu. urcovani stari a pomeru pohlavi u okouna (Pen a fluviatilis 
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1906 Der Karpfen. Geschichte. Naturgeschichte und wirtschaftliche Bedeutung 
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I ESZCZYNSKA. W. AND I . BINIAKOWSK] 

1967 Chow sazana w osrodku zarybieniowym PZW Otorowo [Breeding of the 
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1962 O sviazi kontinentov Evropy i Ameriki [About the unity of European and 
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1957 Uber Hungerformen beim Brassen. Der Fischwirt, vol. 8. 

MAAR. A 

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1966 Carp breeding in drainage tt ater. Bamidgeh, vol. 1 8. pp. 5 1-54. 

MCCRJMMON. H. R. 

1968 Carp in Canada. Bull. Fish. Res. Bd. Can., no. 165. pp. 1-93. 

MISIK. V. 

1958 Biometrika dunajskeho kapra (Cyprinus carpio carpio L.) z dunajskeho 
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MISIK. V. AND V. TUCA 

1965 Posudzovanie extericru dunajskeho kapra so zretel'om na vyber plemen- 
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31 



MORCINLK, P. 

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1948 On the relation between salinity contents of the water and living condition 
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1924 Compendium of Roman History; Res gestae divi Augusti, with an Hnglish 
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pei ikan, o. 

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1881- Pommerellisches Urkundenbuch. 2 vols. Danzig, Westpreussischer Ge- 
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PI inius, s. c. 

1635 Historiae naturalis. I.ugduni Batavorum. 

popova, o. A. 

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1967 Ochrana zivocichov v praxi. K vyhlaskam o ochrane vol'ne zijucich zivo- 
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1961 Vergleichende Untersuchungen iiber den Wildkarpfen der Donau und den 
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RUETIMEYER, L. 

1860 Untcrsuchung der Thierreste aus den Pfahlbauten der Schweiz. Zurich, 
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SEGERSTRALE, C. 

1932 Uber die jahrlichen Zuwachszonen der Schuppen und Beziehungen zwischen 
Sommertemperatur und Zuwachs bei Abramis brama sowie einigen andcren 
Cypriniden in Sud-Finland 1911-1930. Acta Zool. Fenn., vol. 13, pp. 1-42. 

STEFFENS, W. 

1958 Der Karpfen. Wittenberg. A. Ziemsen. 90 pp. 

1959 Wachstum und Kbrperform ungarischer Wildkarpfen in deutschen Teichen. 
Dt. Fisch Ztg, Radebeul, vol. 6. pp. 2 1 3-2 1 8. 

1964 Vergleichende anatomisch-physiologischc Untersuchungen an Wild -und 
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800. 

32 



STEFFENS. W. 

1967 Das Domestikationsproblem heim Karpfcn (Cyprinus carpio L.). Verh. 
int. Verein. theor. angew. l.imnol.. vol., 16. pp. 1441-1448. 

STRABO. A. 

1917- The Geography of Strabo. with an English translation by H. L. Jones. 
1933 Based in part on the unfinished version of J. R. S. Sterrett. 8 vols. London, 
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1600 Opisanie porzadku stawowego y przestrog niektorych domowego gospo- 
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mri mienski, o. 

1573 O sprawie. sypaniu. wymierzaniu i rybieniu stawow [On management, 
construction, measuring and stocking of ponds]. F. Kucharzewski, 1897, 
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tvcitus. p. c. 

1836 The Works of Cornelius Tacitus, translated by A. Murphy. Philadelphia, 
T. Wardle. 742 pp. 

TALBOT. L.M., W. J. A. PAYNE. H. P. LEDGER. I . D. VERDCOURT AND M. H. TALBOT 

1965 The meat production potential of wild animals in Africa; a review of 
biological knowledge. Commonwealth Bureau of Animal Breeding and 
Genetics Technical Communication, no. 16, Farnham Royal, Bucks., Eng., 
Commonwealth Agricultural Bureau. 42 pp. 

TAMURA. T. 

1961 Carp cultivation in Japan. /;; Borgstrom, G., ed. Fish as Food, vol. 1. 

Production, biochemistry and microbiology. New York, Academic Press, 
pp. 103-120. 

TAYLOR. C. R. 

1970 Dehydration and heat: effects on temperature regulations of East African 
ungulates. Am. J. Physiol., vol. 219, no. 4, pp. 1 136-1139. 

1972 Ranching arid lands: Physiology of wild and domestic ungulates in the 
desert. Botswana Notes and Records, sp. ed. no. 1, Proc. Conf. on Sustained 
Production from Semi-Arid Areas. Oct. 1971. Gaberone, pp. 167-192. 
[Mimeographed] 

TCHEN, S. C. 

1956 A history of the domestication and the factors of the varietal formation 
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322. 

7HIFNEMANN, A. 

1950 Verbreitungsgeschichte der Susswassertierwelt Europas. Die Binnenge- 
wasser Band 18. Stuttgart. F. Schweizerbartsche. 809 pp. 

TUCA, v. 

1958 Studium genetickej fexteriero\ej ) cistoty dunajskeho kapra [Study of 
genetical (exterior) purity of the Danube carp]. lab. Fish. Res. Final Rept., 
Bratislava. Typescript. [In Slovak] 

VALCANOVER. F. 

1960 All the paintings of Titian. Hawthorn Books, Inc., New York. Part 2. 
Plate 127-128. 

33 



VARRO, M. T. 

1912 Varro on Farming. M. Terenti Varronis Rerum Rusticarum libri tres, 
translated by L. Storr-Best. London. G. Bell. 375 pp. 

VOLF, J. 

1965 Kralik. Svetem zvirat. Sv. 3. Domaci zvirata [The rabbit. In Hanzak, J., 
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Animals.] Prague. SNDK. [In Czech] 

VOOREN, C. M. 

1972 Ecological aspects of the introduction of fish species into natural habitats 
in Europe, with special reference to the Netherlands. J. Fish Biol., vol. 4, 
no. 4, pp. 565-583. 

WUNDER, W. 

1949 Fortschrittliche Karpfenteichwirtschaft. Stuttgart. 

ZAUNICK, R. 

1925 Tritt der Karpfen schon im Diluvium Norddeutschlands auf ? Mitteilg. d. 
Fischereivereine Brandenburg-Pommern etc., vol. 17, pp. 80-83. 



34 



P e 3 K) m e 

H.ih ycTanoB.ieHHH HCTopiiMecKoro HanaJia pa3Be^eHHH 
Kapna 6bi.in ncno.ib30BaHbi HCTopuMecKne, 3ooreorpa- 
ipnMeckiie, Mopcpo.ioniMecKHe h cpH3HO.iornMecKHe CBe- 
aeHHH. ripeAKii ahkoto Kapna (ca3ana) BepoHTHO 6e- 
p>T CBoe HaMa.io b Cpe^Heft Ashh h noTOM ecTecTBeH- 
hwm nyTeM pacnpocTpaHHioTCH Ha boctok, b KnTafi h 
Ha 3anaa, ao pewn /XyHafi. Ha ocHOBaHHH HCTopnMecKnx 
AaHHbix mo>kho npcancuaraTb, mto pnMJiHHe nepBbie 
Haqa.iH pa3Be^eHne ca3aHa H3 peKH JXyaai'i. H 3Ta Tpa- 
ahuhh Bbuep>KHBaHHH ca3aHa b piscinae npoAOJUKa- 
.iacb b MOHacTbipax b CpeAHHe BeKa. BbwocjiHBOCTb ah- 
Koro Kapna H3 peKH JlyHafl npoAeMOHCTpnpoBaHa Ha 
onbue ro.icuoBaHHH 3toh pbi6bi. Pa3BeAeHHe Kapna 
b KHTae BHaMane 6h\no He3aBncnMbiM ot noAo6Hofi 
jenTe.ibHOCTH EBponbi. yooK^aromHe AOKa3aTejibCTBa 
VKa3biBaK)T Ha to, mto ahkhh Kapn peKH JXynaPi hbjih- 
eTCH npe^KOM mhothx coBpeMeHHbix KyjibTHBnpoBaH- 

HblX BHAOB Kapna. 3TO HB.TfleTCH Ba>KHbIM cpaKTopoM 

b no.ib3y coxpaHeHHH stoh pbi6bi jinn AaJibHei'tuiero 
ynoTpeS.ieHHH. 

nocKo.ibKy jo.ie^HHKOBbie ocTaTKH Kapna b c(paft- 
Hbix nodpoHKax He npn3HaK)T H3-3a hctomhocth hx 
jaTbi, nosTOMv npeAno.iaraioT, mto ahkhh Kapn npo- 
hcxoaht H3 CpeAHen A3HH, oTKyAa oh pacnpocTpa- 
hh.ich ecTecTBeHHbiM nyTeM Ha boctok, b KHTaft h 
CMe>KHbie KpaH h Ha 3anaA, b nocjieAHioK) noc/iejieA- 
hhkobvk) anoxy, ao caMoro Jlyuan. B AOKa3aTe;ibCTBO 
Toro, mto 3to He npoH3onjJio paHbine npnBOAHTcn 
cpaKT, mto b n.iencToneHOByio 3noxy nocre oTxoAa 

KOHTHHeHTa.lbHblX .leAHHKOB C. carpio He nOHBHJICH 
BMeCTe C flpyrHMH BHAaMH pbl6, KOTOpbie npOHHK- 

jih b CKaHj.HHaBHK), Ha BpHTaHCKne OcTpoBa h b Ce- 
BepHyio AMepuKv. Ecjih 6bi b stot nepHOA Kapn no- 

HBH.1CH B BOCTOMHOH A3HH, TO OH B03MO/KHO nOHBHJI- 

ca 6bi h b CeaepHOH AMepnKe, ecTecTBeHHbiM nyTeM, 
BMecTe co myKOH h ffpyrHMH npecHOBOAHbiMH pbi6a- 

MH, KOTOpbie npOHHKJH TVRSL Mepe3 BepHHTOB npOJIHB. 

B aoKa3aTe.ibCTBo ecTecTBeHHoro nonB;ieHHH ah- 
koto Kapna b J\ynae B3hth c.icayioniHe cpaKTbi: pa3- 
HbiMH HMeHaMH Ha3biBa.in Kapna MecTHi.ie /Khtcjih; 

OAHHM H TeM >Ke KCIbTHHCKHM HMeHeM Ha3bIBa.7IH 3TV 

pi,ioy BHe ee ecTecTBeHHbix 3anajHbix rpamm npe6bi- 
BaHHH; CBeaeHHH H3 CpeAneBeKOBbix vieTonncen o pac- 
npocTpaHeHHH Kapna; H3 coBpeMeHHbix nccjeAOBaHHii 
o ahkom Kapne, >KHBymeM b peKe JXynan. ZlncpcpepeH- 
unanHH ahkoto Kapna b AanewoM npouiJiOM b CpeA- 
Hefi A3HH H IIOHB.ieHHe HeH3MeHHBIJJHXCfl riOOAHHOKHX 

rpynn 3toh pbi6bi b boctomhux h 3anaAHbix panoHax 

HaBOAMT Ha MblClb TOM, MTO MeCTOM IipoHCXO>KAeHHH 

35 



Kapna /xo.iyKHa 6i,iTb Cpcahhh A31151. 

Oi\a.e;ibHaH rviaBa nocBnmaeTCH onbiTy, iicc.iejyio- 
meMy cnoco6HOCTb ahkoto Kapna peKH JXynaPi Bbi- 
>KHTb b ycjioBnnx rojiona. 3tot onbiT no^TBen>Kj.aeT 
BbiHoc.iHBocTb 3TOM pi>i6bi h o6-bHCHHeT noqeMy Kapna 

p33BO^HJIH JIK),HH AJIH nHUIH. 

HaKOHeu, n3 3annceft Phm.ihh n H3 BbiKonaHHbix 
npe^MeTOB HCKyccTBa, n3o6pa>KaK)iHHx Kapna, nanpa- 
ujHBaeTCH rnnoTe3a o tom, mto BnepBbie 3Ta pbi6a 
nonajia b pbi6Hi>ie ca^.KH Phmjihh H3 Tex MecT, me npo- 
xOAHJia fluTapHan ,H,opora — caivian rjiaBnan Aopora 
Tex BpeMen, nepeceKaroman peKy Zlynan. 3to 6bijih 
MecTa peryjinpHo HaBOAHennbix ji\tob b ycTbe peKH 
MopaBbi, okojio iipeBnero KapHyHTyiwa h 3aMKa XXe- 
BHHa, rjie o6biKHOBeHHO CTan Kapna MeTajiH HKpy. IlaT- 

PHUHII, KOMaHAHpbl PHMCKHX JieTHOHOB, CTOHBUIHX B 

9thx MecTax, 3aBe3^n Kapna b Phm, KaK aniiKypufi- 
ckhh ^e.nnKaTec noTOiuy, mto 3to 6biJia ojwa H3 ca- 
mwx BbmocjiHBbix bhaob pi>i6, KOTopaa CMoraa nepe- 
>KHTb npnMnTnBHyio TexHHKy TpaHcnopTa h pa3Be,n,eHHe 
pi»i6i.i Tex BpeMeH. Ilocjie yna^Ka h pa3pyuieHiiji Phm- 
ckoh HMnepnn TexHHKa pa3Be^ennH Kapna b pumckhx 
puGHbix ca^Kax He 6biJia 3a6poujeHa. 3to neno npo- 
nojixiajiu xpncTHHHCKne MHccnoHepbi n MOHaxn, Hy>K- 
AaioiunecH b 3anacax pi>i6i>i bo BpeMH pe,iHrH03in>ix 
nocTOB. Mncciionepbi 11 Monaxn 3HajiH 113 jiaTHHCKHX 
3annceii, a TaK>Ke H3 onbiTa k»khi>ix cTpaH, OTKyna 
ohh BbiiiiJin o tom, mto i<apn no^xo^HT /uiH pa3Be,ae- 

IIHH H OHH 3aHHMaJIHCb 3THM fleJIOM B MeCTaX CBOdO 

hoboto noce^eHHH. C kohhom Cpe/mnx BeKOB pa3Be- 
Aenne Kapna 6i»ijio y>Ke pyTHHoft ne TOJibKO b MOHa- 

CTbipHX, HO H B MaCTHblX HMeilHHX. 

HcTopnH pa3Be^.enHH Kapna b EBpone, na i iiiiiaio- 
uiancn jxuKHM KapnoM H3 peKH .Hynan (Cyprinus carpio 
carpio), ac/ihtch na nHTb nepHO^OB: 1 nepno/i, b 

kotopom nepBbiMH ikhuuih pa3Bo^nTb Kapna Phm- 
;hihc, Ha 3ana,n. ot peKH JXyuavi, b nepBOM n flo neT- 
Beproro ctojicthh Hauieii spti; 2 - nepnoA cnopa/ui- 
necKoro iianajia pa3Be,ueHHfl Kapna b iihtom h inec- 
tom cTOJieTHJix; 3 - nepnoA MaccoBoro pa3Be,aeHini 
Kapna 11 cTpcMJiemiH k pa3Be,n.eHHK> stoh pw6bi b 6ojib- 
iiiom Ko:inuecTBe b cneu.na.ibno 3aBCAeHHbix npyaax b 
ce^bMOM 11 no TpimaaHaToe CTO.ieTiin; 4 — nepnoA 
iiinpoKO pacnpocTpaiiemioro pa3BeAeHiui Kapna n ciy- 
MaiHioe ce^eKHHoinipoBaiuic b ucTbipnaAuaTOM 11 no 
niecTHaAHaToe ctojicthh; 5 nepnoA ycujieHiioro 

pa3BeAeiiHH, ucieycTpeMJieHHan ce.ieKunn 11 nanajio 
pa3BeAeniiH Kapna b CeBepnoii AMcpiiKe, ABCTpa.iHH, 
AajibneM Boctokc n b AcppiiKe c ceivuiaAuaToro cto- 
jieTHH. Bo3mo>kho b KuTac pa3Be,aeHne Kapna naMa- 
;h)ci» ii3 MecTHoro noABH^a C. c. haematopterus 500 

36 



.ieT paHbiue, ho He 38Bhchmo ot Ebpoiii.i. 

3;j.ocb noa i iepKiiiuieTCH ca&roe Baxuioe cbomctbo, 
no KOTopoMv aiiKHii Kapn peKH /Iynai'i OTJiHiaeTCfl 
ot Tenepeimiero Ky/ibTiiBiipoBannoro Kapna n flaeTCfl 
ccbunca Ha pajiiTo.'ihiioc cxoactbo MexcAy pa3Be,aenn- 
om Kapna n npiipymiBaHiieM KpoJiHKa. Taione o6pa- 
maeTcn BuiiMannc na noTpe6nocTb coxpanenHn noc- 
.lejmix ahkhx craii Kapna b npe^ropbHx peKH Jlyuati 
H npejoxpanenne jiiKoro npe^Ka KWibTiiBiipoBaHHoro 
wapna b ero ecTecTBennou cpeae. II caMoe rjiaBHoe, 
noBTopeHHe cyji.b6bi Bbiwepujero Typa, npe^Ka Kpyn- 
hoto poraToro CKOTa, He aojdkho 6biTb AonycTHMo. 
KaK KpynnbiH poraTbift ckot, TaK ii Ky.ibTHBHpoBaii- 

HblH Kapn B AHKOM COCTOHHHH HB.IHeTCH pbl60H n.10- 

xoro KawecTBa. 3to, bo3.\io>kho, h HB.iHeTCH npHMHHOH 
-roro, mo Kapn cra;i He nony.inpubiM b AiuepHKe noc- 
."ie ero hhtpciykuhh. 



37 



ISBN: 0-8885